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Development of the population of the European Fallow , dama dama (Linnaeus, 1758), in

(Mammalia: Cervidae)

Zeynel Arslangündoğdu, Max Kasparek, Halil Sarıbaşak, M. Süleyman Kaçar, Osman Yöntem, M. Tuğrul Şahin

Abstract. In order to conserve the last autochthonous population of the , Dama dama dama (Linnaeus, 1758), the Turkish Government began a breeding programme at Düzlerçamı near Antalya in 1966. The programme began with 7 and the numbers con- tinuously increased until the mid-1980s, when they reached over 500 animals. However, the population then collapsed until the year 2000 and did not recover. Today it comprises less than 130 individuals. The reasons for the population collapse are not fully understood but are thought to be a combination of several factors related to increasing human pressure such as urbanisation, recreational activities, and . The population is much below the carrying capacity of the area. Attempts to re-introduce Fallow Deer into other areas of Turkey have not been successful but should be further considered as an option to minimise the risk of extinction, as at present the entire gene pool of the Turkish autochthonous population is concentrated at Düzlerçamı.

Key words. Antalya, Düzlerçamı, reintroduction, breeding programme, .

Introduction The Fallow Deer is represented by two different subspecies: the , Dama dama mesopotamica (Brooke, 1875) and the European Fallow Deer, Dama dama dama (Linnaeus, 1758). Some authors separate them at the level. They can be distinguished by their different sizes (Persian Deer are larger) and by differences in the and the tail colouring (HALTENROTH 1959). Recent genetic studies using mitochondrial DNA have con- firmed that D. d. mesopotamica is strongly differentiated from D. d. dama (MASSETI et al. 2006, 2008). Both forms were originally neighbours but have been isolated by geographic barriers in the east and south/southeast of Turkey. The range of the Persian Fallow Deer extended from Iran over Iraq to Syria and Palestine, and included some parts of southern and south-eastern Turkey. European Fallow Deer existed north and west of this area (HEIDE- MANN 1987). The Persian Fallow Deer, which was previously abundant throughout western Asia, is currently listed in the IUCN Red List as endangered (RABIEI 2008). A report from 1994 estimated the wild population in Iran to be no more than 15 animals (HEIDEMANN 1994), and today the global population is thought to comprise less that 250 mature individu- als (RABIEI 2008). As a result of introductions by the Phoenicians, Romans, and others, however,

Zoology in the Middle East 49, 2010: 3-12. ISSN 0939-7140 © Kasparek Verlag, Heidelberg 4 Zoology in the Middle East 49, 2010

the European Fallow Deer is widespread and abundant in and is listed by IUCN as of least concern. Its original range is unclear, but current knowledge suggests that there were post-glacial refuges of the species in Turkey and southern and south-eastern Europe (e.g. HEIDEMANN 1976, CHAPMAN & CHAPMAN 1975, 1980, MASSETI 1999). In its Turkish native range, it was once widespread and not rare (HEIDEMANN 1976, UERPMANN 1981), but it is now under serious threat (MASSETI & MERTZANIDOU 2008). The range extended along the Mediterranean coastline, south-western , the Aegean region, some parts of the Mar- mara region and the southern parts of eastern Anatolia (HUŞ 1974, HEIDEMANN 1976, KUMERLOEVE 1975, TURAN 1984). The exact borderline between Persian and European Fallow Deer in Turkey is not known. A marked reduction in the number of Fallow Deer in Anatolia took place when the lowland forests were progressively transformed into agricul- tural land and hunting increased (HALTENROTH 1959, HEIDEMANN 1976, KUMERLOEVE 1975, HEIDEMANN 1987, TURAN 1987). The species became extinct in the Marmara and Aegean regions at the turn of 20th century. Only populations on the islands around Ayvalık, at Adaköy near and in the Bay of Gökova appear to have survived longer (MASSETI & MERTZANIDOU 2008). The process of declining numbers and even of extinctions in some parts of the Mediterranean region became worse after the 1950s (CHAPMAN & CHAPMAN 1975, HEIDEMANN 1976, TURAN 1987). As a response to the declining numbers, the Turkish Government launched in the 1960s a breeding and conservation programme at Düzlerçamı in Antalya province, where Turkey’s only undoubtedly natural wild population of Fallow Deer was living. The effort to preserve the last remaining autochthonous population still continues. The aim of this study is to com- pile all available information on the population living in and around the enclosed breeding station, to assess the species’ current population status in Turkey, and to determine the over- all success of the breeding programme. For this purpose we used information from published sources, unpublished internal reports and our own field surveys.

The Düzlerçamı Reserve An area of 1,750 ha, in which 2 males, 2 females and 3 fawns were living, was designated in 1964 as the Fallow Deer Conservation Area (HUŞ 1964, TURAN 1966). It was later enlarged to 11,432 ha and became a Fallow Deer and Wild Conservation Area. In 1970, the area was further enlarged to 14,300 ha and the populations of both species began to recover dramatically thanks to strict conservation measures. In 1987, the area finally received wildlife , and by that time had reached 34,000 ha (SARIBAŞAK et al. 2005). Adult Fallow Deer studs reared in Düzlerçamı were then sent to other similar stations in various parts of the country, such as the islands around Ayvalık (Ayvalık-Adalar), Gökova and Adaköy in Muğla province and Pos- Çatalan in Adana province (TURAN 1984, ÜSTAY 1990, MASSETI 1999, MASSETI 2007). When food shortages began, the administration decided to relocate Fallow Deer in 2002. Within the species’ natural habitat, a new 521 ha expanse, 430 ha of which was fenced, housed a new breed- ing station and 64 deer were re-located (SARIBAŞAK et al. 2005). In 2005, the area was designated the Düzlerçamı Wildlife Development Area and now comprises 28,973 ha. Düzlerçamı Wildlife Development Area is located between 36°55’–37°03’ northern latitude and 30°28’–30°34’ eastern longitude. Elevation of the area ranges from 250 to 350 m. The study area comprises 7,500 ha of forest and 600 ha of open agricultural land at the Düzlerçamı Wildlife Development Area (Fig. 1). Tree composition consists of dense stands of Calabrian Pine (Pinus brutia) with dense shrubs in the lower forest structure, as well as open areas and privately owned agricultural fields. The flora consists of a total of 426 plant species, 33 of which are endemic to

Mammalia 5

Fig. 1. Extension of Düzlerçamı Wildlife Development Area and the study area in Antalya province, Turkey.

the area (SARIBAŞAK et al. 2007). The area reflects typical Mediterranean climate characteristics, namely hot and dry summers, and mild and humid winters. The area includes a 122 km forest road network, together with strategically installed fire strips and buffers. In summer, an average of 50 motor vehicles visits the area daily. Although poaching is prohibited, Fallow Deer is hunted using the practice of stand hunting. There is an approximate population of 5,485 people living in the nearby villages of Yukarı Karaman, Nebiler, Duraliler, Aşağı Karaman, Doyran, Çığlık and Bayat Bademleri, and these villagers graze their livestock inside the conservation area. More than 1,000 grazing animals from the villages of Yukarı Kara- man and Doyran are guarded by 25 to 30 shepherd dogs.

Counting techniques To carry out a census of the numbers of Fallow Deer, drive, concealed waiting and indirect enu- meration techniques (CAUGHLEY & SINCLAIR 1994, OĞURLU 2003) were used in collaboration with the Antalya Natural Conservation and National Parks Service from 2005–2008. To identify the difficulties the animals face during breeding, we investigated behavioural patterns, common locations, the pressure of poaching, and how the animals utilize the designated areas. Given the fact that the growth ratio was 70%, and utilising natural birth and death rates, a theo- retical population development (TPD), initialized in 1966, was generated in order to formulate hypotheses about the natural course of population evolution. According to the calculations of

6 Zoology in the Middle East 49, 2010

Table 1. The population recovery rates of Fallow Deer in the Antalya/Düzlerçamı region. In parentheses: inside the enclosure, outside the enclosure. Total area in ha. Number = number of Fallow Deer.

Year Conservation areas Area Number Reference 1966 Fallow Deer Conservation 1,750 7 TURAN 1966, 1969a Area (3 ♂, 3 ♀, 1 young) 1967 Fallow Deer- 7,750 9 TURAN 1967–75, Conservation Area (7 adults, 2 young) 1969a 1968 Fallow Deer-Wild Goat 11,432 16 TURAN 1969a Conservation Area (9 adults & 3 young, 4 from the environs) 1969 " " 22 TURAN 1967–75, (16 adults, 6 young) 1969a 1970 " 14,300 19 TURAN 1967–75 1975 " " 65-70 TURAN 1967–75 " " 60-80 HEIDEMANN 1976 1977 " " 1 ♂ from Thrace & 2 ANONYMOUS 1978 pregant females from the environs brought to breeding station c.1983 " " >500 TURAN 1984 1987 Wildlife Conservation Area 34,000 400 ANONYMOUS 1990 1980s “ “ 400 ZENGINGÖNÜL 1987 1990 " " 300-400 ANONYMOUS 1990 1993 " " 250 ANONYMOUS 1990 1994 " " >537(37 + >500) MASSETI 2007 1996 " " 200 KAÇAR 2002 2000 " " 60-80 KAÇAR 2002 " " 100 (40 + 60) MASSETI 2007 2001 " " 79 (51+28) SARIBAŞAK et al. 2005 2002 " " 85 (57+28) SARIBAŞAK et al. 2005 2003 " " 92 (64+28) SARIBAŞAK et al. 2005 2004 " " 105 (78+27) SARIBAŞAK et al. 2005 150 in total MASSETI 2007 2005 Wildlife Development 28,973 77 (53+24) This study Area 102-103 (52-53+<50) MASSETI 2007 2006 " " 20-31 (20-31+X) ALBAYRAK & ŞAHIN 2006* 90 (60+30) This study 2007 " " 95 (65+30) This study 2008 " " 97 (67+30) This study 2009 " " 126 (96+30) This study

MEHLITZ (1989) and PETRAK (2006), the carrying capacity of a 100 ha area is 4 animals with a gender rate of 1:1. For the calculation of Fallow Deer numbers in the breeding station, the num- ber of females in spring is taken into account, and the annual average population increase is computed at about 70 to 77% (UECKERMANN 1972, MEHLITZ 1989). Mammalia 7

Results The development of the population size of Fallow Deer based on various sources during the 43 years from 1966 to 2009 is shown in Table 1 and Fig. 2. The year 1966 was taken as the benchmark for calculating the theoretical population development during this 43-year-long period. An initial population of 2 males, 2 females and 3 fawns at a gender ratio of 1:1 was the basis for these calculations (Fig. 2). The current Fallow Deer habitat comprises an 8,100 ha area within the total 28,973 ha of the Wildlife Development Area. Additionally, there is an unused area of 10,378 ha that could be used by these animals in the future. The carrying capacity of the 8,100 ha in the core area is 324 animals, but the 739 animals in the 10,378 ha of the development area are included. The habitat actually used by Fallow Deer extends over an area of 22,000 ha. This area in- cludes 3,532 ha of rocky terrain in which the ancient city of Termessos was once located and which constitutes a particularly important habitat for Fallow Deer. A further 10,378 ha is considered to be a potential habitat for the future of the Fallow Deer, but no has been reported to live in this area since the beginning of the study. The remaining area includes 3,760 ha occupied primarily by houses that are used on a seasonal basis, 107.9 ha of agricul- tural and open fields, including olive and orange groves, 5.6 ha used for fresh water fisher- ies, and 27.6 ha that was fenced off out of a total of 4,330 ha that are heavily utilized by Fallow Deer. It was observed that the population of Fallow Deer is likely to go foraging before dusk and very early in the morning. They prefer the green pastures from early in the spring until the summer, when their feeding habits shift from grass to the leaves and shoots of maquis ele- ments during the summer months. They also feed on the leaves of coniferous trees during these periods. During the autumn months, faecal examination results showed that they feed on acorn, carob, berries and olive kernels. Males eat large quantities of these kernels, espe- cially in the mating season. Finally, in the winter, it was found that they feed on grasses, fallen oak leaves and acorn kernels. Other large that use the habitat as a common area in the Wildlife Development Area include , Vulpes vulpes (Linnaeus, 1758), Wild Cat, Felis silvestris Schreber, 1777, Caracal, Caracal caracal (Schreber, 1776), Eurasian , Sus scrofa Linnaeus, 1758, and Wild Goat, aegagrus Erxleben, 1777 (SARIBAŞAK et al. 2005, DE MARINIS & MASSETI 2009). There are indications that the Anatolian Leopard, Panthera pardus tul- liana (Valenciennes, 1856) survived in the area at least until the early 1990s (ULLRICH & RIFFEL 1993) and the Striped Hyaena, Hyaena hyaena (Linnaeus, 1758), until the early 1980s (KASPAREK et al. 2004).

Discussion Düzlerçamı, Antalya, is habitat for the last remaining autochthonous population of Fallow Deer. Through the establishment of a conservation area and the implementation of strict conservation measures, the population has increased from less than ten animals in the mid- 1960s to over 500 in the mid-1980s. However, the numbers then started to decline and the current population comprises less than 130 animals. The reason for this decline is not fully understood. Both a rapid urbanisation around the Wildlife Conservation Area and a rela- tively large forest fire affecting 2,100 ha forest area in 1997 possibly contributed to the shrinkage of the Fallow Deer habitat; as a result, Fallow Deer foraging and sheltering

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800 Theoretical population development Real population development 700

600

500

Number of Fallow Deer 400

300

200

100

0 1966 1968 1970 1972 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 2008

Fig. 2. Theoretical population development (TPD) and real population development (RPD) of Fallow Deer in the Düzlerçamı Wildlife Development Area.

resources were adversely affected (KAÇAR 2002). 13 Fallow Deer became sick and died in 1999 because of the quality of the fodder used at the breeding station. Also, poaching could not be fully controlled and it is still common in the region. Fallow Deer hunting is part of the nomadic culture of the villagers at Yukarı Karaman and Aşağı Karaman, and control mecha- nisms have failed so far mainly because the inhabitants of these two villages are all related. Caracal and Wild Boar may have some negative impact, especially on the offspring of Fal- low Deer. However, the number of predators has decreased in recent decades (Anatolian Leopard and Striped Hyaena have disappeared) and there are no indications that the numbers of shepherd dogs and stray dogs have increased. An important factor, however, may be poi- soning by local people: fugitive hunters who were caught in the breeding station were found to be poisoning Fallow Deer. In 2005, 25 animals died inside the breeding station for this reason. The motives of these people are not fully understood. The pressure on the area is increasing, especially with the growing population and more people visiting the conservation area. The number of settlements around the conservation area is increasing, and industrial areas in the west and south of Antalya are constantly ex- panding. According to figures provided by the local Statistical Office, the human population in the province of Antalya increased between 1990 and 2000 by 4.4% in the urban centres and 4% in the suburbs. In the conservation area there are now a trout restaurant and related facilities, beekeeping activities and private properties, and there are also farms and gardens as well as government tree plantations which are surrounded with fences and thus restrict Fallow Deer movements. This places continuously increasing pressure and stress on the deer population. The increased human impact is also evident in the form of forest fires, poaching, grazing of domesticated animals and beekeeping (SARIBAŞAK et al. 2005). Mammalia 9

The carrying capacity of the 8,100 ha core area was calculated as 324 animals, or as 739 animals if the 10,378 ha development area is taken as a basis. This is less than SARIBAŞAK et al. (2005) suggested; they believed that 800 animals could live in the 8100 ha core area (8-10 animals per 100 ha). The theoretical population recovery course progressed successfully until 1984 but did not continue after 1987 when the numbers started to decrease. These fluc- tuations cannot be related to the carrying capacity of the area. The numbers of Fallow Deer at Düzlerçamı and other places are unfortunately not always very precise and are sometimes even contradictory. Table 1 gives different figures for some years according to different sources. There were, for example, approximately 1300 Fallow Deer said to be living in Turkey in 1986 (TARHAN 1986), while another source gave a total of 700 animals in all conservation areas and breeding stations combined for 1987 (TURAN 1987). At about the same time, 600 animals were reported from Düzlerçamı (TARHAN 1987) and more than 500 Fallow Deer were given by ERKAN (1987). These figures seem to be over-estimates and are not regarded as reliable by us. They show how difficult it is to obtain reliable figures and how necessary it is to conduct long-term monitoring with standardised methods. Several attempts have been made to reintroduce Fallow Deer into other parts of the coun- try with the help of animals from the Düzlerçamı Breeding Station. Fallow Deer were apparently surviving in the forests of the Bay of Gökova (Muğla prov- th ince) until the middle of the 20 century, but already NEBIOĞLU (1968) noted that this popu- lation had ceased to exist. A breeding station was established by the then Ministry of For- estry at Akyaka in the Bay of Gökova (Muğla province) at some time in the 1980s (see also TURAN 1984) and a few animals were translocated from Düzlerçamı. In the 1980s and 1990s, on average approximately six animals were living in the enclosure (B. SUSEVEN, pers. comm.). MASSETI (2007) saw two adult males and three females there in 1994. The small population was reproducing but all the young animals died (B. SUSEVEN, pers. comm.). The breeding station, which also supported Wild Goat, was closed in the early 2000s and the remaining animals were brought to Düzlerçamı. Today it is no longer operational (M. KAS- PAREK). A viable population of Fallow Deer was found up to the middle of the 20th century to the north of Adana. When TURAN (1969b) went to the forests of Çatalan, Pos Sarıçam and Sırk- ıntı in 1969, he met local hunters who gave plausible accounts that in their whole lives they had hunted altogether 50 or even 100 Fallow Deer. At the time of his visit, only 10-12 ani- mals remained. Upon TURAN’s initiative, a 9,000 ha “Fallow Deer Conservation Area” was established (TURAN 1969b, 1984). One pair of Fallow Deer that was transferred from Düzlerçamı to Çatalan in 1986 was lost in 1991 due to inadequate care (İNAÇ 1994). The conservation area included a 15 ha enclosure (Karaisalı-Çatalan Alageyik Yetiştirme Alanı). According to information provided by the Provincial Directorate for Environment and For- estry in Adana in March 2010, Fallow Deer have existed in the area for many years. MASSETI (2007) saw Fallow Deer in a fenced area near Mersin in 1994 but the exact loca- tion and the origin of these animals are not known. Finally, three Fallow Deer, including an adult male, were sent from Düzlerçamı to İstanbul (Akfıratlar, Tuzla) in 1994 (MASSETI 2007). The fact that a male Fallow Deer was captured in Thrace in 1977 (and subsequently brought to Düzlerçamı) may indicate that a small population existed there at that time. NE- BIOĞLU (1968) also mentioned areas around Manavgat and Alanya, both in the Antalya prov- ince, where probably 50-60 animals had survived. A further 5-10 animals were reported to him to have survived between Burdur and Antalya (NEBIOĞLU 1968). 10 Zoology in the Middle East 49, 2010

The idea of introducing Fallow Deer into areas other than Düzlerçamı is in principal largely supported by us. Even if the species does not succeed in establishing additional stable populations, these areas may serve as genetic reserves. As the Düzlerçamı population is at present the only surviving autochthonous population, it is at a high risk of extinction. It was discussed whether diseases introduced by domestic may affect the Düzlerçamı popula- tion and may cause a population decline. In such a situation, animals surviving outside Düzlerçamı may preserve the gene pool. The fact that a male Fallow Deer was brought in 1977 from European Turkey (Thrace) to Düzlerçamı for reproduction is evidence that the genetic constitution of the Düzlerçamı population may already have suffered and is not as pure as had been thought. Fallow Deer is also brought to Turkish zoos from abroad from time to time. The zoo of İzmir, for example, imported two male and two female Fallow Deer from Bulgaria in March 2010, the zoo of Gaziantep has 21 Fallow Deer with unknown origin and the zoo of Antalya seven (four male, three female), which they received from Ankara and Kahraman Maraş. Two female Fallow Deer were brought from Belgium and the Netherlands, respectively, to a private farm in Hatay in 2006. Although these animals are normally kept in captivity, a mixture with local animals, i.e. interbreeding of different genetic lineages can finally not be ruled out. Recent studies on the genetic constitution of Fallow Deer have shown that there is a high genetic diversity within the species and even close populations such as the one on are ge- netically very distinct (MASSETI et al. 2006, 2008). An introduction of animals from outside Turkey could therefore destroy the genetic constitution of the autochthonous Turkish popula- tion and must be avoided by all means. HEIDEMANN (1987) called the import of non- domestic breeds to Turkey a “zoological crime”. Proposals by the Directorate of Nature Conservation and National Parks to supplement local Fallow Deer stocks with those from other countries are not supported by us.

Acknowledgements. We would like to thank Serhat YILDIRIM, Provincial Directorate of Environment and Forestry of Antalya, Bahar Heike SUSEVEN, Akyaka, and Ziver TUNA, İstanbul University Forestry Faculty, for assisting us in preparing this paper. We would further like to thank all local staff and game wardens who take care of the Fallow Deer at Düzlerçamı and who helped in conducting the counts.

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Authors’ addresses: Dr Zeynel Arslangündoğdu, Istanbul University, Faculty of Forestry, Department of Forest Entomology and Protection, 34473 Bahçeköy-Sarıyer, Istanbul, Turkey. – Dr Max Kasparek, Mönchhofstr. 16, 69120 Heidelberg, Germany. – Halil Sarıbaşak & M. Süleyman Kaçar, Southwest Anatolia Forest Research Institute, Antalya, Turkey. – Osman Yöntem & M. Tuğrul Şahin, Provincial Directorate of Environment and Forest of Antalya, Antalya, Turkey. – Email: [email protected].